Debate over ‘selfish gene’ theory heats up | Genetic Literacy Project Debate over ‘selfish gene’ theory heats up Printer Friendly (Credit: Flickr/micahb37) The concept of the “selfish gene” has been around for more than three decades. First coined by evolutionary biologist Richard Dawkins in his 1976 book, The Selfish Gene, the term describes sequences of DNA that spread by forming additional copies of itself within the genome and make no specific contribution to the reproductive success of the organism in which it is found. This notion became very popular, in large part, because it captured the idea that all living things, including humans, primarily act to maximize the number of their descendants. It also served as an addendum to Darwin’s Theory of Evolution, helping reinforce the validity of the theory of the survival of the fittest. From the beginning, however, this idea has been questioned by many other scientists, researchers and journalists who scoff at the idea that humans are motivated by their genes and not by their own free will. Now, after David Dobbs wrote about the need to challenge the Selfish Gene Theory, in aeon magazine , new life is being breathed into this decades-long argument.

In the provocatively titled “Die, Selfish Gene, Die,” Dobbs writes about the notion of rapid gene expression—the idea that the genes within any given organism can be re-read or re-expressed to make the organism better suited for survival. Evidence of this theory, Dobbs writes, demands that the scientific community revisit the idea of the selfish gene. “The selfish gene is one of the most successful science metaphors ever invented. Unfortunately, it’s wrong,” he writes. Dobbs explains: For a century, the primary account of evolution has emphasised the gene’s role as architect: a gene creates a trait that either proves advantageous or not, and is thus selected for, changing a species for the better, or not. Thus, a genetic blueprint creates traits and drives evolution. This gene-centric view, as it is known, is the one you learnt in high school. It’s the one you hear or read of in almost every popular account of how genes create traits and drive evolution. It comes from Gregor Mendel and the work he did with peas in the 1860s. Since then, and especially over the past 50 years, this notion has assumed the weight, solidity, and rootedness of an immovable object.

But a number of biologists argue that we need to replace this gene-centric view with one that more heavily emphasises the role of gene expression — that we need to see the gene less as an architect and more as a member of a collaborative remodelling and maintenance crew. Almost immediately after Dobbs’s article was published, it was met with some sharp criticism. “However good Dobbs’s other writing may have been, this is a dire piece: one that is loaded with misinformation, wrong information, misleading information, and unsupported speculation,” wrote science writer Jerry Coyne . “None of it even comes close to deposing the value of the ‘selfish gene’ metaphor, and I’m not saying that just because I’m friends with Richard Dawkins. According to Coyne, Dobbs makes several mistakes: First, he wants to claim that the metaphor of the selfish gene is wrong. Second, he wants to show that it’s wrong because new understanding of gene regulation—how genes turn on and off during development—render the selfish gene metaphor passé.

Finally, he claims that a new theory, that of “genetic accommodation,” relegates much of conventional evolutionary theory to the dustbin, for the new theory deposes the centrality of the gene in favor of the centrality of the environment and its non-genetic effects on development. There were numerous other sharply critical responses to the Dobbs piece. “While I admire Jerry’s take-down of Dobbs, I’m not sure that he (Jerry Coyne) fully appreciates these other criticisms of the selfish gene,” writes science writer Laurence A. Moran, adding: I think we should refer to modern evolutionary theory as “modern evolutionary theory” in order to make sure we’re not talking about “Darwinism,” “neo-Darwinism,” or the hardened version of the “Modern Synthesis.” Modern evolutionary theory includes an important role for random genetic drift, Neutral Theory, and population genetics. We could clarify a lot of discussion if we stopped talking about extending “Darwinism” or extending the Modern Synthesis or proclaiming once again that the selfish gene has died. In fact, the selfish gene has died, it died almost thirty years ago but most people don’t know that. RIP.

Dobbs responded to the criticism in a post on his own website, admitting that he may have “muddled” the message in his original piece: Most crucially, I seem to have not made clear that my challenge was less to a technical account of nature than to a metaphor and story used to describe those technicalities. To put it another way: I apparently did not make clear that “Die, Selfish Gene, Die” is a story less about how genetics and evolution work than about the stories we tell about how genetics and evolution work—and, most crucially, about how those stories about nature percolate out beyond academia and into the minds of the lay public. Dobbs’s article, and the pushback it received, is a good example of how scientific theories are seldom unanimously agreed upon or remain static. But that is not a bad thing. It is the constant challenge that helps drive science forward.

Dobbs writes of his critics, “Despite discord they have engaged under the noble assumption that drives all the best science and all the best writing: That, absent evidence otherwise, we are all here trying to tell true and constructive stories about a nature gorgeously and maddeningly complicated.” Additional Resources: The 100 best nonfiction books: No 10 – The Selfish Gene by Richard Dawkins | Books | The Guardian 100 Best Nonfiction Books of All Time The 100 best nonfiction books: No 10 – The Selfish Gene by Richard Dawkins An intoxicating renewal of evolutionary theory that coined the idea of the meme and paved the way for Professor Dawkins’s later, more polemical works ‘Infectious brio’: Richard Dawkins in 2013.

Photograph: Andy Hall for the Observer 100 Best Nonfiction Books of All Time The 100 best nonfiction books: No 10 – The Selfish Gene by Richard Dawkins An intoxicating renewal of evolutionary theory that coined the idea of the meme and paved the way for Professor Dawkins’s later, more polemical works Monday 4 April 2016 00.45 EDT Last modified on Wednesday 1 June 2016 10.18 EDT Share on Messenger Close What is man, and what are we for? Remarkably, it was not until Charles Darwin published On the Origin of Species in 1859 that anyone, in our history, had thought methodically to address the reason for our existence. Darwin’s answer to this simple question was to show that every earthly species – chimps or humans, lizards or fungus – had evolved over about 3bn years by the process known as natural selection. But then, after the furious controversy surrounding that publication, Darwin’s celebrated theory fell into neglect and misuse.

A hundred years later, in the heady, innovative atmosphere of the 1960s, a new generation of young and ambitious evolutionary biologists found themselves confronted with a rare opportunity: the rediscovery and renewal of evolutionary theory. Enter Richard Dawkins, a young Oxford zoologist who had been born, and partly raised, in Africa. Following some notable pioneers such as WD Hamilton and GC Williams, Dawkins pulled together many disparate strands of thought about the nature of natural selection, and organised them into a conceptual framework with far-reaching implications for our understanding of Darwin’s ideas. He called it The Selfish Gene, a title he later considered to contain an unconscious echo of Oscar Wilde’s The Selfish Giant. Dawkins was convinced that an amplified and developed version of neo-Darwinism “could make everything about life fall into place, in the heart as well as in the brain”. His book would extol, he wrote, a “gene’s-eye view of evolution”. It was Dawkins’s simple, but profound, proposition that “the fundamental unit of selection, and therefore of self-interest, is not the species, nor the group, nor even, strictly, the individual.

It is the gene, the unit of heredity.” He acknowledged that this might sound “at first like an extreme view” but proceeded to explore all the major themes of social theory in the light of this idea, conducting his survey in a highly readable and entertaining way. With chapters such as “Genesmanship”, “Battle of the Sexes”, and “Nice Guys Finish First”, he tackled concepts of altruism and selfishness, the evolution of aggressive behaviour, kinship theory, sex ratio theory, reciprocal altruism, deceit, and the natural selection of sex differences. In hindsight, it seems appropriate that The Selfish Gene should have been published soon after The Joy of Sex and The Female Eunuch. You can call me a big bad wolf but not a bore, says Richard Dawkins Read more From his first page, Dawkins unfolds an exhilarating and combative narrative of the gene’s-eye view of life with infectious brio. The Selfish Gene, he declares, should be read “almost as though it were science fiction.

It is designed to appeal to the imagination.” Part of the book’s compulsion derives from Dawkins’s appealing certainty that he is exploring a scientific world in which “we are survival machines – robot vehicles blindly programmed to preserve the selfish molecules known as genes”. This insight, he reports, is “a truth which still fills me with astonishment”. Much of this book’s appeal lies in its author’s barely suppressed excitement, prose that bubbles over with the intoxication of a brilliant new approach. He buttonholes his readers; he dazzles with paradox and provocation. In an introduction to a later edition of The Selfish Gene , Dawkins describes its gestation: it was a book written in extremis (the power cuts and industrial strife of the early 1970s) and, as he says, “in a fever of excitement”. For the young author, it was, in hindsight, “one of those mysterious periods in which new ideas are hovering in the air”. So zeitgeisty was it that, from first publication, the reception of The Selfish Gene was highly favourable. Initially, it was not seen as a controversial book, Dawkins wrote later.

“Its reputation for contentiousness took years to grow.” Eventually it would become regarded as a work of extreme radicalism. But, he goes on, “over the very same years as the book’s reputation for extremism has escalated, its actual content has seemed less and less extreme”. Richard Dawkins: 'I don't think I am strident or aggressive' Read more This is undeniable: while The Selfish Gene grew out of orthodox neo-Darwinian ideas, it actually expressed Darwinism in a way that Darwin himself might have welcomed. Rather than focus on the individual organism, it looked at nature from the perspective of the gene. It was, claimed Dawkins, “a different way of seeing, not a different theory”. It also addressed itself to “three imaginary readers”: the generalist, the expert and the student. This was a high-low cohort that swiftly propelled it on to bestseller lists worldwide. Moreover, in keeping with the temper of the times, The Selfish Gene announced itself, from the first, as “a book about animal behaviour”, arguing that “we, like all other animals, are (survival) machines created by our genes”. For Dawkins, “we” did not mean just people.

He wanted his description to embrace all animals, plants, bacteria and viruses. “The total number of survival machines on Earth,” he wrote, “is very difficult to count.” Even the total number of species is unknown, he conceded. “Taking just insects alone, the number of individual insects may be a million million million.” Dawkins – in a style that would recur in later polemical books such as The God Delusion – was never less than comprehensive in his ambitions. Here, he used “survival machine” rather than “animal” because he wanted to encompass all plants and humans, too. His argument should apply, he said, to any and all “evolved beings”. Orthodox neo-Darwinian he might be, but in chapter 11, he coined an idea about cultural transmission that quickly went viral within the global intellectual community: the meme, or replicator, a unit of imitation.

Examples of memes include tunes, idea, catchphrases, clothes fashions, ways of making pots or of building arches: “Just as genes propagate themselves in the gene pool by leaping from body to body via sperms or eggs, so memes propagate themselves in the meme pool by leaping from brain to brain via a process which, in the broad sense, can be called imitation.” Shortly after this analysis, Dawkins characterised “God” as a meme. Thus, pages 192-193 of The Selfish Gene might be said to encapsulate most of Richard Dawkins’s brilliant career, in which the theory of evolution came to offer such a satisfying and complete explanation for the complexity of life on Earth that there could no longer be a place for the possibility of God’s design. A signature sentence “A monkey is a machine that preserves genes up trees, a fish is a machine that preserves genes in the water; there is even a small worm that preserves genes in German beer mats. DNA works in mysterious ways.” Three to compare EO Wilson: On Human Nature (1978) WD Hamilton: Narrow Roads of Gene Land, Vol I (1996) Why are genes selfish? Edward H.

Hagen, Institute for Theoretical Biology, Berlin Why are genes selfish? Richard Dawkins wrote a very popular book called the Selfish Gene that explained, for a popular audience, many of the exciting new theories and discoveries being made in evolutionary biology in the 1960�s and 70�s. The metaphor Dawkins chose, the selfish gene, was an extremely powerful metaphor, so powerful that it has often overshadowed the science itself! The controversies that swirl around EP are often tightly bound up with Dawkins� metaphor. If our genes are selfish, are we all, deep down, unalterably selfish ourselves? Why did Dawkins chose this metaphor, what does it really mean, and what are its implications for EP and human nature? Simplifying greatly for the sake of the argument, there is a molecule, called a nucleotide, that comes in four different types, A, C, G, & T. Large numbers of these nucleotides can be linked together in a linear strand to make a much bigger molecule called DNA. Schematically, DNA looks like this: ACGTGCCT�etc. Human DNA consists of about 3 billion nucleotides chained together.

Simplifying greatly again, small sections of the DNA strand called �genes�, which are usually several hundred to several thousand nucleotides long, are able to create a different type of long, linear molecule called a protein. Proteins are a kind of plastic�technically, a polymer�that, like the DNA chain, are made up of a small number of different molecular building blocks called amino acids (there are 20 different amino acid building blocks ). Just as the different chemical structure of the plastic in a plastic bag versus the chemical structure of the plastic in dental floss versus the chemical structure of the plastic in bullet-proof vests gives these different materials very different properties, the exact sequence of the different amino acids in a particular protein determines its biological properties. Different amino acid sequences give different proteins very different properties. To summarize, the sequence of nucleotides in small sections of our DNA (called genes) determines the sequence of amino acids of proteins created by the genes, and these amino acid sequences determine the proteins� biological properties.

Although scientists are still debating the exact number, DNA contains somewhere between 30,000 and 60,000 different genes, and can therefore create between 30,000 and 60,000 different proteins, each with unique properties. As I mentioned, proteins are a kind of plastic, so our DNA functions, in part, to create a large number of different plastics with different properties. These highly specialized plastics with very special biological properties are what our bodies are made of. DNA is a kind of lumber yard that provides, among other things, a large number of plastic building materials for making organisms. Simplifying once more (this time by ignoring sex), individuals pass on an exact copy of their DNA chain to their offspring: if my body is made up of a particular set of plastics, because my offspring has an exact copy of my DNA, my offspring�s body will be make up of exactly the same set of plastics, and so it will be exactly like me.

Occasionally, however, one of the molecules in the DNA chain (i.e., one of the A, C, G, or T nucleotides), can become mutated (altered) by cosmic radiation, environmental toxins, etc.; these mutagens turn one type of nucleotide into another type of nucleotide (e.g., an A turns into G). If I pass on this mutated DNA, where only one of the 3 billion nucleotides is different from my own, then my offspring will be made of proteins that are almost exactly like mine, except for the protein which was made by the mutated section of DNA (the mutated gene), which will be a different protein with different properties. My offspring will not be exactly like me�he will be said to have a different phenotype (body type). Imagine that the gene that was mutated was the gene that made the plastic forming the lens of the eye. This plastic has a very special property: it is almost completely transparent. Most proteins, like those forming your skin, muscles, hair, etc., are not transparent.

Because my offspring has a mutated form of the lens gene, there are now two types of genes in the population that make the lens protein: the normal version, possessed by most individuals (which I will call Tnormal, for normal transparency) and the mutated version possessed by my offspring. Different forms of the same gene are called alleles. Because it is far easier to make something worse than it is to make it better, most of the time when the gene producing the lens protein is mutated (and this happens very rarely), the altered lens protein produced by the mutated gene will not be as transparent as the original version (so I will label this allele Tlow, for low transparency). My offspring will therefore not be able to see as well as other members of his species who have the Tnormal allele, and thus normal versions of the lens protein. Let�s assume that my offspring�s lens protein is only slightly less transparent than the normal version. He will be able to live his life and have offspring. The population will therefore contain a mix of lens alleles; most of the population will have Tnormal, but some will have Tlow.

Because those with the Tlow allele will not be able to see quite as well as other members of their species possessing Tnormal, on average they will not have as many offspring. Perhaps they notice prey slightly less often, and thus not have quite as much food, or perhaps they fail to notice predators slightly more often, and will therefore be killed and eaten at a slightly higher frequency. Over many, many generations, the fraction of individuals with the Tlow allele will decrease relative to those with the Tnormal allele simply because individuals possessing the Tlow allele produce, on average, fewer offspring. We say that the Tlow allele producing the less transparent lens protein is selected against, and that the frequency of this allele decreases with time. Notice that, if the total population size remains constant, that the decrease in frequency of the Tlow allele results in an increase in the frequency of the Tnormal allele. Imagine another mutation of the normal lens allele, creating a third allele (Tsuper) that produces a super transparent lens protein. The population now contains three alleles, Tlow, Tnormal, and Tsuper.

Individuals possessing the Tsuper allele detect prey, on average, at slightly higher frequencies, and thus have more food, and they more frequently detect predators and therefore are eaten slightly less frequently. As a consequence, on average, they have more offspring than individuals possessing Tnormal. Over many, many generations, the frequency of the Tsuper allele will increase in the population, whereas the frequencies of the Tnormal and Tlow alleles will decrease, and perhaps disappear altogether. This is called evolution by natural selection: the frequencies of the three alleles have changed as a consequence of their reproductive effects. Over time, a population will acquire alleles that produce proteins that better solve critical reproductive problems, and lose alleles that produce proteins that less effectively solve these problems. There is widespread agreement that evolution by natural selection is responsible for the origins of the sophisticated organs and tissues like hearts, lung, livers, etc., that enable organisms to reproduce. Because, in a population of a given size, the increase in the frequency of Tsuper must decrease the frequencies of the other alleles, biologists began saying that different alleles were �competing�.

(Usually, but not always, alleles increase their frequency by causing individuals possessing them to produce, on average, more offspring.) Dawkins, highlighting the iron-clad logic that alleles increase their frequency in the population if they cause more copies of themselves to be made relative to other alleles, and that by increasing their own frequency, they decrease the frequency of the alternative (competing) alleles, termed genes �selfish�. Alleles increase their frequency at the expense of other alleles. Copyright 1999-2002 Edward H. Hagen In retrospect: The selfish gene : Nature : Nature Research Matt Ridley reassesses Richard Dawkins's pivotal reframing of evolution, 40 years on. The Selfish Gene Richard Dawkins Oxford University Press: 1976. Terry Smith/The LIFE Images Collection/Getty Richard Dawkins in 1976, around the time he published his first best-selling book. Books about science tend to fall into two categories: those that explain it to lay people in the hope of cultivating a wide readership, and those that try to persuade fellow scientists to support a new theory, usually with equations. Books that achieve both — changing science and reaching the public — are rare.

Charles Darwin's On the Origin of Species (1859) was one. The Selfish Gene by Richard Dawkins is another. From the moment of its publication 40 years ago, it has been a sparkling best-seller and a scientific game-changer. The gene-centred view of evolution that Dawkins championed and crystallized is now central both to evolutionary theorizing and to lay commentaries on natural history such as wildlife documentaries. A bird or a bee risks its life and health to bring its offspring into the world not to help itself, and certainly not to help its species — the prevailing, lazy thinking of the 1960s, even among luminaries of evolution such as Julian Huxley and Konrad Lorenz — but (unconsciously) so that its genes go on. Genes that cause birds and bees to breed survive at the expense of other genes. No other explanation makes sense, although some insist that there are other ways to tell the story (see K. Laland et al. Nature 514, 161–164; 2014 ).

What stood out was Dawkins's radical insistence that the digital information in a gene is effectively immortal and must be the primary unit of selection. No other unit shows such persistence — not chromosomes, not individuals, not groups and not species. These are ephemeral vehicles for genes, just as rowing boats are vehicles for the talents of rowers (his analogy). As an example of how the book changed science as well as explained it, a throwaway remark by Dawkins led to an entirely new theory in genomics. In the third chapter, he raised the then-new conundrum of excess DNA. It was dawning on molecular biologists that humans possessed 30–50 times more DNA than they needed for protein-coding genes; some species, such as lungfish, had even more. About the usefulness of this “apparently surplus DNA”, Dawkins wrote that “from the point of view of the selfish genes themselves there is no paradox. The true 'purpose' of DNA is to survive, no more and no less.

The simplest way to explain the surplus DNA is to suppose that it is a parasite.” “Dawkins's ideas helped to explain what was going on inside genomes long before DNA sequencing became routine.” Four years later, two pairs of scientists published papers in Nature formally setting out this theory of “selfish DNA”, and acknowledged Dawkins as their inspiration ( L. E. Orgel and F. H. C. Crick Nature 284, 604–607 (1980) ; W. F. Doolittle and C. Sapienza Nature 284, 601–603; 1980 ). Since then, Dawkins's speculation has been borne out by the discovery that much surplus DNA consists of reverse transcriptase — a viral enzyme whose job is to spread copies of itself — or simplified versions of transposons dependent on it. Thus, Dawkins's ideas helped to explain what was going on inside genomes, as well as between individuals, even though the book was written long before DNA sequencing became routine. The complexity of the structure of the gene itself has since grown enormously, with the discovery of introns, control sequences, RNA genes, alternative splicing and more.

But the essential idea of a gene as a unit of heritable information remains, and Dawkins's synthesis stands to this day. On The Selfish Gene's 30th anniversary, many of Dawkins's admirers, including writer Philip Pullman and cognitive scientist Steven Pinker, contributed essays to the book Richard Dawkins (Oxford University Press, 2006) edited by his former students Alan Grafen and Mark Ridley (no relation of mine). In this Festschrift, the philosopher Daniel Dennett argued that the book was not just science, but “philosophy at its best”. In my contribution, I pointed out that the success of the book had spawned a gold rush for popular-science writers, as publishers began offering large advances in the hope of finding the next Selfish Gene. James Gleick's Chaos (Abacus, 1988), Stephen Hawking's A Brief History of Time (Bantam, 1988) and Pinker's The Language Instinct (William Morrow, 1994) were among the nuggets mined before the boom petered out.

Although his book brimmed with original thoughts, Dawkins was quick to acknowledge that he was building on the discoveries and insights of others, notably the evolutionary theorists William Hamilton, George Williams, John Maynard Smith and Robert Trivers. They were equally quick to appreciate that he had done something more than explain their ideas. Trivers wrote the foreword, and Maynard Smith narrated a television documentary about the book soon after it was published. Williams said in an interview that Dawkins's book had “advanced things a lot further than mine did” (see go.nature.com/21j1mt ); Hamilton wrote that The Selfish Gene “succeeds in the seemingly impossible task of using simple, untechnical English to present some rather recondite and quasi-mathematical themes of recent evolutionary thought” in a way that would “surprise and refresh even many research biologists” ( W. D. Hamilton Science 196, 757–759; 1977 ). The Fayetteville Observer/AP/PA Dawkins speaking at an atheist event in 2012.

As a first-year undergraduate in the zoology department at the University of Oxford, UK, where Dawkins was about to teach me computing and animal behaviour, I found the book exhilarating and bewildering. Until then, my teachers had helpfully divided the world into right ideas and wrong ones. But here was a writer turning some settled science upside down and inviting me to join him on a journey to discover a truth that seemed to him “stranger than fiction”. Was he right or wrong? I was being shown the arguments, not the answers. The origin of The Selfish Gene is intriguing. Dawkins revealed in the first volume of his memoirs, An Appetite for Wonder (Bantam, 2013; see E. Scott Nature 501, 163; 2013 ), that the idea of selfish genes was born ten years before the book was published. In 1966, the Dutch biologist Niko Tinbergen asked Dawkins, then a research assistant with a new doctorate in animal behaviour, to give some lectures in his stead.

Inspired by Hamilton, Dawkins wrote in his notes (reproduced in An Appetite for Wonder): “Genes are in a sense immortal. They pass through the generations, reshuffling themselves each time they pass from parent to offspring ... Natural selection will favour those genes which build themselves a body which is most likely to succeed in handing down safely to the next generation a large number of replicas of those genes ... our basic expectation on the basis of the orthodox, neo-Darwinian theory of evolution is that Genes will be 'selfish'.” Dawkins began writing the book in 1973, and resumed it in 1975 while on sabbatical. At the suggestion of Desmond Morris, the zoologist and author of The Naked Ape (Jonathan Cape, 1967), Dawkins showed some draft chapters to Tom Maschler of Jonathan Cape, who strongly urged that the title be changed to 'The Immortal Gene'. Today, Dawkins regrets not taking the advice.

It might have short-circuited the endless arguments, so beloved of his critics and so redolent of the intentional stance (in which we tend to impute mental abilities to unconscious things, from thunderstorms to plants), about whether selfishness need be conscious. It might even have avoided the common misconception that Dawkins was advocating individual selfishness. In the end, it was Michael Rodgers of Oxford University Press who enthusiastically published The Selfish Gene, after demanding “I must have that book!” when he saw early draft chapters. It was an immediate success, garnering more than 100 reviews, mostly positive. Dawkins went on to write books that were better in certain ways. The Extended Phenotype was more groundbreaking, The Blind Watchmaker more persuasive, Climbing Mount Improbable more logical, River out of Eden and Unweaving the Rainbow more lyrical, The Ancestor's Tale more encyclopaedic, The God Delusion more controversial. But they were all variations on the themes he so eloquently and adventurously set out in The Selfish Gene. George C. Williams, Leading Evolutionary Theorist, Dies at 83 - The New York Times The New York Times Science |George C.

Williams, 83, Theorist on Evolution, Dies Search George C. Williams, an evolutionary biologist who helped shape modern theories of natural selection, died Wednesday at his home in South Setauket on Long Island, near Stony Brook University, where he taught for 30 years. He was 83. The cause was Parkinson’s disease, said his wife, Doris Williams. Dr. Williams played a leading role in establishing the now-prevailing, though not unanimous, view among evolutionary biologists that natural selection works at the level of the gene and the individual and not for the benefit of the group or species. He is “widely regarded by peers in his field as one of the most influential and incisive evolutionary theorists of the 20th century,” said Douglas Futuyma, a colleague and the author of a leading textbook on evolution. Dr. Williams laid out his ideas in 1966 in his book “Adaptation and Natural Selection.” In it, he seized on and clarified an issue at the heart of evolutionary theory: whether natural selection works by favoring the survival of elements as small as a single gene or its components, or by favoring those as large as a whole species.

He did not rule out the possibility that selection could work at many levels. But he concluded that in practice this almost never happens, and that selection should be understood as acting at the level of the individual gene. Advertisement Continue reading the main story In explaining an organism’s genetic adaptation to its environment, he wrote, “one should assume the adequacy of the simplest form of natural selection” — that of variation in the genes — “unless the evidence clearly shows that this theory does not suffice.” The importance of Dr. Williams’s book was immediately recognized by evolutionary biologists, and his ideas reached a wider audience when they were described by Richard Dawkins in his book “The Selfish Gene” (1976). Those ideas have continued to draw attention because group selection still has influential advocates. In highly social organisms like ants and people, behaviors like altruism, morality and even religion can be more directly explained if selection is assumed to favor the survival of groups. Dr. Williams had a remarkably open turn of mind, which allowed him always to consider alternatives to his own ideas. David Sloan Wilson, a leading advocate of group selection, recalled in an interview that as a graduate student he once strode into Dr.

Williams’s office saying he would change the professor’s mind about group selection. “His response was to offer me a postdoctoral position on the spot,” Dr. Wilson said. Photo Credit Michael Shavel Dr. Wilson did not take the position but remained close to Dr. Williams, though the two continued to differ. One matter of dispute was whether a human being and the microbes in the gut and the skin could together be considered a superorganism created by group selection. Dr. Williams did not believe in superorganisms. (Nonetheless, when Dr. Wilson came to visit him one day, Dr. Williams had taped to his door a hand-lettered sign saying, “Superorganisms welcome here.”) Dr. Williams’s interests extended to questions that evolution seemed not to answer well: Why should a woman forfeit her chance of having more babies by entering menopause? Why do people grow old and die when nature should find it far easier to maintain a body than to build one? An important article he wrote in 1957 on the nature of senescence led to a collaboration with Dr. Randolph Nesse, a psychiatrist at the University of Michigan.

Together they developed the concept of Darwinian medicine, described in the 1995 book “Why We Get Sick.” There the authors offered Darwinian explanations for questions like why appetite decreases during a fever or why children loathe dark green vegetables. Dr. Williams pursued his ideas even to results that he found disturbing. “He concluded that anything shaped by natural selection was inevitably evil because selfish organisms outproduced those that weren’t selfish,” Dr. Nesse said. Advertisement Continue reading the main story Dr. Williams acknowledged that people had moral instincts that overcome evil. But he had no patience with biologists who argue that these instincts could have been brought into being by natural selection. “I account for morality as an accidental capability produced, in its boundless stupidity, by a biological process that is normally opposed to the expression of such a capability,” Dr. Williams wrote starkly in 1988. In the field of evolutionary theory, “George was probably the most influential author in the 1960s,” said William Provine, a historian of evolution at Cornell University. But by choosing important subjects, Dr. Williams remained relevant.

His ideas were approachable because he wrote in clear, simple prose and largely without the use of mathematics, an almost obligatory tool for most evolutionary biologists today. Dr. Williams joined the State University of New York at Stony Brook (now Stony Brook University) in 1960 and worked there until his retirement in 1990. In addition to his wife, who is also a biologist, he is survived by a son, Jacques; three daughters, Sibyl Costell, Phoebe Anderson and Judith Pitsiokos; and nine grandchildren. Though a major expositor of evolutionary theory, Dr. Williams was always aware that his explanations were a work in progress and that they might in principle be superseded by better ones. Evolutionary theory, as stated by its great 20th-century masters Ronald Fisher, J. B. S. Haldane and Sewall Wright, “may not, in any absolute sense, represent the truth,” Dr.

Williams wrote at the conclusion of his book on adaptation, “but I am convinced that it is the light and the way.” A version of this article appears in print on September 14, 2010, on Page A27 of the New York edition with the headline: George Williams, 83, Theorist on Evolution, Dies. Order Reprints | Today's Paper | Subscribe Selfless Genes: A New Revolution in Biology | The Huffington Post Selfless Genes: A New Revolution in Biology 04/02/2012 09:53 am ET | Updated Jun 02, 2012 Jonathan Gottschall Author, 'The Storytelling Animal: How Stories Make Us Human' In 2006, I attended the annual meeting of the Human Behavior and Evolution Society in Austin, Texas and listened to E. O. Wilson -- the founder of sociobiology and one of the greatest biologists since Darwin -- deliver the keynote address. Wilson, about 75 years old at the time, didn't come to Austin to play the grand old man. He came to pick a fight.

When it was over, a buzz of excitement filled the room: "Did he really just call selfish gene theory 'a monumental mistake?'" Yes he did. And he repeats the charge in his fascinating and controversial new book, The Social Conquest of the Earth . In the late 1960s and early 1970s evolutionary biologists celebrated a fundamental breakthrough. William Hamilton's inclusive fitness theory (aka selfish gene theory ) indicated that organisms are narrowly "designed" to spread copies of their own genes, whether those genes are located in their own bodies or in the bodies of their relatives. Hamilton's work seemed to show exactly how evolution worked, and also how it didn't work. Group selection -- the idea that competition between groups of organisms shapes genomes -- was declared dead. In effect, this defined altruism -- real and authentic selflessness -- out of existence. On a planet ruled by selfish genes, "altruism" was just masked selfishness. The biologist Michael Ghiselin expressed this beautifully, "Scratch an altruist and watch a hypocrite bleed." The Selfish People Let's run a quick thought experiment to see how biologists reached this conclusion.

Imagine that long before people spread out of Africa there was a tribe called The Selfless People who lived on an isolated island off the African coast. The Selfless People were instinctive altruists, and their world was an Eden. But then there was the fall. A genetic mutation produced a single tribesman who was naughty, not nice. He had a simple prime directive encoded in his DNA: Look out for number one (and for your kin folk -- and perhaps your allies, but only if you're sure they'll reciprocate). The tribesman had selfish genes. The biological definition of selfishness: behavior that promotes one's own reproductive success. The biological definition of selflessness: behavior that promotes someone else's reproductive success at the expense of one's own. The single tribesman with selfish genes would -- by definition -- leave more descendants behind, eventually crowding altruistic traits out of the gene pool. The Selfless People would become The Selfish People. The big 1960s breakthrough was simply this: selfish genes beat selfless genes; they beat them bloody; they beat them every single time. The Selfless People For a generation, this logic ruled evolutionary biology, and especially evolutionary psychology.

But most of the biologists who were responsible for the 1960s breakthrough have gradually backed off their positions (the major hold out is Richard Dawkins, who writes that the "great heresy" of group selection "really is wrong"). Our thought experiment played out the way it had to. But that's because it had one unrealistic component: The Selfless People live in total isolation from all other tribes. And this isolation makes all the difference. Now, let's run the thought experiment more realistically. This time the Selfless People live in a rich African valley that they share with another tribe. The two tribes frequently come into conflict over hunting grounds, old grudges, and insults barked across the river. And so the tribes fight, as tribes of men have always fought. Imagine that the valley's second tribe is comprised mainly of selfish actors. Other factors held equal, who wins: the tribe of self-sacrificing altruists or the tribe where every warrior is looking out for number one? Won't it be the Selfless People? Won't the Selfless People tend to dominate selfish tribes in most competitive situations? And, as a result, won't selfless genes proliferate? Charles Darwin thought so.

In The Descent of Man, Darwin ran his own thought experiment, pitting selfless against selfish tribes: It must not be forgotten that although a high standard of morality gives but a slight or no advantage to each individual man and his children over the other men of the same tribe, yet that an advancement in the standard of morality and an increase in the number of well-endowed men will certainly give an immense advantage to one tribe over another. There can be no doubt that a tribe including many members who, from possessing in a high degree the spirit of patriotism, fidelity, obedience, courage, and sympathy, were always ready to give aid to each other and to sacrifice themselves for the common good, would be victorious over other tribes; and this would be natural selection. Here, Darwin describes how group-level competition can produce authentic altruism. In The Social Conquest of the Earth, Wilson updates Darwin's case, mainly drawing on his expertise as an entomologist. Wilson argues that the incredible levels of cooperation and altruism within ant colonies testify to millions of years of vicious conflict between colonies.

Darwin and Wilson agree: no matter the species, if you have intense and sustained group-level conflict, selfless genes beat selfish genes; they beat them bloody; they beat them every single time. Of course, it would be a great distortion to suggest that people are -- like ants -- selfless all of the time. But the vision of rigid selfishness that arose from biology's rejection of group selection was an equally great distortion. The real picture is more complex. Natural selection occurs at the level of groups and individuals. Between-group competition favors selfless genes while competition inside groups favors selfish genes. As Wilson and a colleague wrote , "Selfishness beats altruism within groups. Altruistic groups beat selfish groups. Everything else is commentary." The Superorganism vs. the Selfish Gene - How Natural Selection Works | HowStuffWorks The Superorganism vs. the Selfish Gene Emanuele Biggi/ Getty Images Evolutionary biologist Richard Dawkins wrote a book called "The Selfish Gene" in the 1970s. Dawkins' book reframed evolution by pointing out that natural selection favors the passing on of genes, not the organism itself. Once an organism has successfully reproduced, natural selection doesn't care what happens after.

This explains why certain strange traits continue to exist -- traits that seem to cause harm to the organism but benefit the genes. In some spider species, the female eats the male after mating. As far as natural selection is concerned, a male spider that dies 30 seconds after mating is just as successful as one that lives a full, rich life. Since the publication of "The Selfish Gene," most biologists agree that Dawkins' ideas explain a great deal about natural selection, but they don't answer everything. One of the main sticking points is altruism. Why do people (and many animal species) do good things for others, even when it offers no direct benefit to themselves? Research has shown that this behavior is instinctive and appears without cultural training in human infants [source: CBC ]. It also appears in some primate species. Why would natural selection favor an instinct to help others? Up Next How Kin Selection Works One theory revolves around kinship. People who are related to you share many of your genes. Helping them could help ensure that some of your genes are passed down. Imagine two families of early humans, both competing for the same food sources. One family has alleles for altruism -- they help each other hunt and share food.

The other family doesn't -- they hunt separately, and each human only eats whatever he can catch. The cooperative group is more likely to achieve reproductive success, passing along the alleles for altruism. Biologists are also exploring a concept known as the superorganism. It's basically an organism made out of many smaller organisms. The model superorganism is the insect colony. In an ant colony, only the queen and a few males will ever pass their genes to the next generation. Thousands of other ants spend their entire lives as workers or drones with absolutely no chance of passing on their genes directly. Yet they work to contribute to the success of the colony. In terms of the "selfish gene," this doesn't make a whole lot of sense. But if you look at an insect colony as a single organism made up of many small parts (the ants), it does. Each ant works to ensure the reproductive success of the colony as a whole. Some scientists think the superorganism concept can be used to explain some aspects of human evolution [source: Wired Science ]. Vestigial and Atavistic Traits All organisms carry traits that no longer confer any real benefit to them in terms of natural selection.

If the trait doesn't harm the organism, then natural selection won't weed it out, so these traits stick around for generations. The result: organs and behaviors that no longer serve their original purpose. These traits are called vestigial. There are many examples in the human body alone. The tailbone is the remnant of an ancestor's tail, and the ability to wiggle your ears is left over from an earlier primate that was able to move his ears around to pinpoint sounds. Plants have vestigial traits as well. Many plants that once reproduced sexually (requiring pollination by insects) evolved the ability to reproduce asexually. They no longer need insects to pollinate them, but they still produce flowers, which were originally needed to entice insects to visit the plant. Sometimes, a mutation causes a vestigial trait to express itself more fully. This is known as an atavism . Humans are sometimes born with small tails. It's fairly common to find whales with hind legs. Sometimes snakes have the equivalent of toenails, even though they don't have toes. Or feet.

Question: Who is the British evolutionary theorist who wrote 'The Selfish Gene'?
Answer: RICHARD DAWKINS